Metamorphosis

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A dragonfly in its final moult, undergoing metamorphosis from itsnymph form to an adult.
Metamorphosis is a biological process by which an animal physically develops after birth or
hatching, involving a conspicuous and relatively abrupt change in the animal's body structure
through cell growth and differentiation.
Some insects, amphibians, molluscs, crustaceans, cnidarians, echinoderms and tunicates under
go metamorphosis, which is usually accompanied by a change of habitat or behavior.
Scientific usage of the term is exclusive, and is not applied to general aspects of cell growth,
including rapid growth spurts. References to "metamorphosis" inmammals are imprecise and
only colloquial, but historically idealist ideas of transformation and monadology, as
in Goethe's Metamorphosis of Plants, influenced the development of ideas of evolution.
Contents
[hide]
1 Etymology
2 Insect metamorphosis
o 2.1 Hormonal control
3 Amphibian metamorphosis
o 3.1 Frogs and toads
o 3.2 Newts
4 Metamorphosis in fish
5 See also
6 References
7 Bibliography
8 External links
Etymology[edit]
The word "metamorphosis" derives from Greek , "transformation,
transforming",
[1]
from - (meta-), "change" and (morphe), "form".
[2]

Insect metamorphosis[edit]

Incomplete metamorphosis in the grasshopper with different instar nymphs
All insects in the Pterygota undergo a marked change in form, texture and physical appearance
or metamorphosis, from immature to adult. These insects either
have hemimetabolous development, and undergo an incomplete or partial metamorphosis,
or holometabolous development, which undergo a complete metamorphosis, including a pupal or
resting stage between the larval and adult forms.
[3]

In hemimetabolous insects, immature stages are called nymphs. Development proceeds in
repeated stages of growth and ecdysis (moulting); these stages are called instars. The juvenile
forms closely resemble adults', but are smaller and lack adult features such as wings and
genitalia. This process is known as "partial" or "incomplete" metamorphosis. The differences
between nymphs in different instars are small, often just differences in body proportions and the
number of segments, although external wing buds will form in later instars.
In holometabolous insects, immature stages are called larvae, and differ markedly from adults.
Insects which undergo holometabolism pass through a larval stage, then enter an inactive state
called pupa, or chrysalis, and finally emerge as adults. This process is called "complete"
metamorphosis. It is theorized that the pupal stage is the evolutionary compaction of all the
nymphal stages of their hemimetabolous ancestors, while the larval stage is an extended, mobile
form of the developing embryo.
[3]

According to recent research, adult Manduca sexta is able to retain the behaviour learned as
a caterpillar.
[4]
Another caterpillar, the Ornate Moth caterpillar, is able to carry toxins that it
acquires from its diet through metamorphosis and into adulthood, where the toxins still serve for
protection against predators.
[5]

Many observations have indicated that programmed cell death plays a considerable role during
physiological processes of multicellular organisms, particularly
during embryogenesis and metamorphosis.
Sequence illustrating complete metamorphosis in cabbage white butterfly, ''Pieris rapae''

larva

pupa

pupa ready to hatch

adult
Hormonal control[edit]
Insect growth and metamorphosis are controlled by hormones synthesized by endocrine
glands near the front of the body. Neurosecretory cells in an insect's brain secrete a hormone,
theprothoracicotropic hormone (PTTH) that activates prothoracic glands, which secrete a second
hormone, usually Ecdysone (an ecdysteroid), that induces ecdysis.
[6]

PTTH also stimulates the corpora allata, a retrocerebral organ, to produce juvenile
hormone (JH), which prevents the development of adult characteristics during ecdysis. In
holometabolous insects, molts between larval instars have a high level of JH, the moult to the
pupal stage has a low level of JH, and the final, or imaginal, molt has no JH present at
all.
[7]
Experiments on Firebugs have also shown how JH can affect the number of nymph instar
stages in hemimetabolous insects.
[8][9]

Amphibian metamorphosis[edit]

Just before metamorphosis, only 24 hours are needed to reach the stage in the next picture

Almost functional common frog with some remains of the gill sac and a not fully developed jaw
In typical amphibian development, eggs are laid in water and larvae are adapted to an aquatic
lifestyle. Frogs, toads, and newts all hatch from the egg as larvae with external gills but it will take
some while for the amphibians to interact outside with pulmonary respiration. Afterwards, newt
larvae start a predatory lifestyle, while tadpoles mostly scrape food off surfaces with their horny
tooth ridges.
Metamorphosis in amphibians is regulated by thyroxin concentration in the blood, which
stimulates metamorphosis, and prolactin, which counteracts its effect. Specific events are
dependent on threshold values for different tissues. Because most embryonic development is
outside the parental body, development is subject to many adaptations due to specific ecological
circumstances. For this reason tadpoles can have horny ridges for teeth, whiskers, and fins.
They also make use of the lateral line organ. After metamorphosis, these organs become
redundant and will be resorbed by controlled cell death, called apoptosis. The amount of
adaptation to specific ecological circumstances is remarkable, with many discoveries still being
made.
Frogs and toads[edit]
With frogs and toads, the external gills of the newly hatched tadpole are covered with a gill sac
after a few days, and lungs are quickly formed. Front legs are formed under the gill sac, and
hindlegs are visible a few days later. Following that there is usually a longer stage during which
the tadpole lives off a vegetarian diet. Tadpoles use a relatively long, spiralshaped gut to digest
that diet.
Rapid changes in the body can then be observed as the lifestyle of the frog changes completely.
The spiralshaped mouth with horny tooth ridges is resorbed together with the spiral gut. The
animal develops a big jaw, and its gills disappear along with its gill sac. Eyes and legs grow
quickly, a tongue is formed, and all this is accompanied by associated changes in the neural
networks (development of stereoscopic vision, loss of the lateral line system, etc.) All this can
happen in about a day, so it is truly a metamorphosis. It isn't until a few days later that the tail is
reabsorbed, due to the higher thyroxin concentrations required for tail resorption.
Newts[edit]

The large external gills of thecrested newt
In newts, there is no true metamorphosis because newt larvae already feed as predators and
continue doing so as adults. Newts' gills are never covered by a gill sac and will be resorbed only
just before the animal leaves the water. Just as in tadpoles, their lungs are functional early, but
newts don't make as much use of them as tadpoles do. Newts often have an aquatic phase in
spring and summer, and a land phase in winter. For adaptation to a water phase, prolactin is the
required hormone, and for adaptation to the land phase, thyroxin. External gills do not return in
subsequent aquatic phases because these are completely absorbed upon leaving the water for
the first time.
Metamorphosis in fish[edit]
Some fish, both bony fish (Osteichthyes) and non-bony fish (Agnatha), undergo metamorphosis.
Fish metamorphosis is typically under strong control by the thyroid hormone.
[citation needed]

Examples among the non-bony fish include the lamprey. Among the bony fish, mechanisms are
varied.
The salmon is diadromous, meaning that it changes from a freshwater to a saltwater lifestyle.
Many species of flatfish begin their life bilaterally symmetrical, with an eye on either side of the
body; but one eye moves to join the other side of the fish - which becomes the upper side - in the
adult form.
The European eel has a number of metamorphoses, from the larval stage to
the leptocephalus stage, then a quick metamorphosis to glass eel at the edge of the continental
shelf (8 days for the Japanese eel), two months at the border of fresh and salt water where the
glass eel undergoes a quick metamorphosis into elver, then a long stage of growth followed by a
more gradual metamorphosis to the migrating phase. In the pre-adult freshwater stage, the eel
also has phenotypic plasticity because fish-eating eels develop very wide mandibles, making the
head look blunt. Leptocephali are common, occurring in all Elopomorpha (Tarpon- and eel-like
fish).
Most other bony fish undergo metamorphosis from embryo to larva (fry) and then to the juvenile
stage during absorption of the yolk sac, because after that phase the individual needs to be able
to feed for itself.
[10]